Table 1. VSV recombinants used as oncolytic agents against cancer.

Recombinant VSV	Virus description	Reference(s)	Designed to improve:
			Oncoselectivity	Safety	Oncotoxicity	VSV survival	Tumour immunity
WT and miscellaneous
WT VSV (‘Rose lab’)	The parental rWT VSV for most VSV-based OVs. The L gene and the N-terminal 49 residues of the N gene are derived from the Mudd-Summers strain, the rest is from the San Juan strain (both Indiana serotype)	Lawson et al. (1995)
VSV-WT-XN2 (or XN1)	Derivative of rWT VSV (‘Rose lab’). Generated using pVSV-XN2 (or pVSV-XN1), a full-length VSV plasmid containing unique XhoI and NheI sites flanked by VSV transcription start and stop signals between G and L genes. pVSV-XN2 (or pVSV-XN1) is commonly used to generate recombinant VSVs encoding an extra gene	Schnell et al. (1996)
WT VSV (‘Wertz lab’)	Alternative rWT VSV. The N, P, M and L genes originate from the San Juan strain; G gene from the Orsay strain (both Indiana serotype). Rarely used in OV studies	Whelan et al. (1995)
VSV-WT-GFP, -RFP, -Luc, -LacZ	WT VSV encoding reporter genes (between G and L) to track virus infection. Based on pVSV-XN2. Toxicity similar to VSV-WT	Fernandez et al. (2002), Wu et al. (2008)
VSV-G/GFP	GFP sequence fused to VSV G gene is inserted between the WT G and L genes (in addition to WT G). Toxicity similar to that of VSV-WT	Dalton & Rose (2001)
VSV-rp30	Derivative of VSV-G/GFP. Generated by positive selection on glioblastoma cells and contains two silent mutations and two missense mutations, one in P and one in L. ‘rp30’ indicates 30 repeated passages	Wollmann et al. (2005)	X	X	X
VSV-p1-GFP, VSV-p1-RFP	VSV expressing GFP or red fluorescent protein (RFP or dsRed) reporter gene at position 1. Attenuated because all VSV genes are moved downward, to positions 2–6. Safe and still effective as an OV	Wollmann et al. (2010)	X	X
VSV-dG-GFP (or RFP) (replication-defective)	Similar to VSV-p1-GFP or VSV-p1-RFP described above, but with the G gene deleted. Cannot generate a second round of infection. Poor ability to kill tumour cells	Wollmann et al. (2010)	X	X
VSV-ΔP, -ΔL, -ΔG (semi-replication-competent)	Each virus cannot replicate alone because of one VSV gene deleted, but when viruses co-infect, they show good replication, safety and oncolysis (especially the combination of VSVΔG/VSVΔL). VSVΔP and VSVΔL contain dsRed in place of the corresponding viral gene. VSVΔG contains GFP gene in place of G	Muik et al. (2012)		X
M mutants
VSV-M51R	The M51R mutation was introduced into M	Kopecky et al. (2001)	X	X
VSV-ΔM51, VSV-ΔM51-GFP, - RFP, -FLuc, -Luc, - LacZ	The ΔM51 mutation was introduced into M. In addition, some recombinants encode a reporter gene between the G and L	Stojdl et al. (2003), Power & Bell (2007), Wu et al. (2008)	X	X
VSV-*Mmut	VSV with a single mutation or combination of mutations at the following M positions: M33A, M51R, V221F and S226R	Hoffmann et al. (2010)	X	X
VSV-M6PY >A4-R34E and other M mutants	The M51R mutation was introduced into the M gene, and, in addition, the mutations in the PSAP motif (residues 37–40) of M	Irie et al. (2007)	X	X
VSV-M(mut)	VSV M residues 52–54 are mutated from DTY to AAA. M(mut) cannot block nuclear mRNA export	Heiber & Barber (2011)	X	X
G mutants
VSV-G5, -G5R, -G6, -G6R	VSV-expressing mutant G with amino acid substitutions at various positions (between residues 100 and 471). Triggers type I IFN secretion as the M51R, but inhibits cellular transcription and host protein translation like WT	Janelle et al. (2011)	X	X
VSV-CT1	The cytoplasmic tail of the G protein was truncated from 29 to 1 aa. Decreased neuropathology, but marginal oncolytic efficacy	Ozduman et al. (2009), Wollmann et al. (2010)	X	X
VSV-CT9-M51	The cytoplasmic tail of VSV-G was reduced from 29 to 9 aa, also has ΔM51 mutation. Attenuated neurotoxicity and good OV abilities	Ozduman et al. (2009), Wollmann et al. (2010)	X	X
Foreign glycoproteins
VSV-DV/F(L289A) (same as rVSV-F)	VSV expressing the NDV fusion protein gene between G and L. The L289A mutation in this protein allows it to induce syncytia alone (without NDV HN protein)	Ebert et al. (2004)	X	X	X
VSV-S-GP	VSV with the native G gene deleted and replaced with a modified glycoprotein protein (GP) from Sindbis virus (SV). Also expressing mouse GM-CSF and GFP (between SV GP and VSV L). The modified GP protein recognizes the Her2 receptor, which is overexpressed on many breast cancer cells	Bergman et al. (2007)	X	X	X
VSV-ΔG-SV5-F	VSV G gene is replaced with the fusogenic simian parainfluenza virus 5 fusion protein (SV5-F) gene	Chang et al. (2010)		X	X
VSV-FAST, VSV-(ΔM51)-FAST	VSV or VSV-MΔ51 expressing the p14 FAST protein of reptilian reovirus (between VSV G and L)	Brown et al. (2009)	X	X	X
VSV-LCMV-GP (replication-defective)	VSV lacking the G gene was pseudotyped with the non-neurotropic glycoprotein of LMCV	Muik et al. (2011)		X
VSV-H/F, -αEGFR, -αFR,-αPSMA (replication-defective)	VSV lacking the G gene was pseudotyped with the MV F and H displaying single-chain antibodies (scFv) specific for epidermal growth factor receptor, folate receptor, or prostate membrane-specific antigen. Retargeted VSV to cells that expressed the targeted receptor	Ayala-Breton et al. (2012)	X	X	X		X
microRNA targets
VSV-let-7wt	The let-7 microRNA targets are inserted into the 3′-UTR of VSV M	Edge et al. (2008)	X	X
VSV-124, -125, -128, -134 (M or L mRNA)	VSV recombinants with neuron-specific microRNA (miR-124, 125, 128 or 134) targets inserted in the 3′-UTR of VSV M or L mRNA	Kelly et al. (2010)	X	X
Cancer suppressors
VSV-mp53, VSV-M(mut)-mp53	VSV [WT or M(mut)] expressing the murine p53 gene. M(mut) has residues 52–54 of the M protein changed from DTY to AAA	Heiber & Barber (2011)	X	X	X		X
Suicide genes
VSV-C : U	VSV expressing E. coli CD/UPRT, catalysing the modification of 5-fluorocytosine into chemotherapeutic 5-FU	Porosnicu et al. (2003)	X		X
VSV-C	VSV-MΔ51 expressing CD/UPRT	Leveille et al. (2011b)	X	X	X
VSV-(MΔ51)-NIS	VSV-MΔ51 expressing the human NIS gene (for ‘radiovirotherapy’ with 131I)	Goel et al. (2007)	X	X	X
VSV-TK	VSV expressing TK; can improve oncolysis if used with non-toxic prodrug ganciclovir	Fernandez et al. (2002)	X		X
Immunomodulation
VSV-mIFNβ, -hIFNβ, VSV-rIFNβ	VSV expressing the murine (m), human (h) or rat (r) IFN-β gene	Obuchi et al. (2003), Jenks et al. (2010)	X	X	X		X
VSV-IL4	VSV expressing IL-4	Fernandez et al. (2002)			X		X
VSV-IL12	VSV expressing IL-12	Shin et al. (2007a)			X		X
VSV-IL23	VSV expressing IL-23. Significantly attenuated in the CNS, but effective OV	Miller et al. (2010)	X	X	X		X
VSV-IL28	VSV expressing IL-28, a member of the type III IFN (IFN-λ) family	Wongthida et al. (2010)	X	X	X		X
VSV-opt.hIL-15	VSV-MΔ51 expressing a highly secreted version of human IL-15	Stephenson et al. (2012)	X	X	X		X
VSV-CD40L	VSV expressing CD40L, a member of the tumour necrosis factor (TNF) family of cell-surface molecules	Galivo et al. (2010)			X		X
VSV-Flt3L	VSV-MΔ51 expressing the soluble form of the human Flt3L, a growth factor activating DCs	Leveille et al. (2011a)	X	X	X		X
VSV/hDCT	VSV-MΔ51 expressing hDCT	Boudreau et al. (2009)	X	X	X		X
VSV-hgp100	VSV expressing hgp100, an altered self-TAA against which tolerance is well-established in C57BL/6 mice	Wongthida et al. (2011b)			X		X
VSV-ova	VSV expressing chicken ovalbumin (for B16ova cancer model)	Diaz et al. (2007)			X		X
VSV-gG	VSV expressing EHV-1 glycoprotein G, a broad-spectrum viral chemokine-binding protein	Altomonte et al. (2008b)			X	X
VSV-UL141	VSV expressing a secreted form of the human cytomegalovirus UL141 protein, known to inhibit the function of NK cells by blocking the ligand of NK cell-activating receptors	Altomonte et al. (2009)			X	X
VSV-(Δ51)-M3	VSV-MΔ51 expressing the murine gammaherpesvirus-68 chemokine-binding protein M3	Wu et al. (2008)	X	X	X	X